Professional Documents
Culture Documents
The Adaptation of Gunda Ulvae To Salinity: Whereas
The Adaptation of Gunda Ulvae To Salinity: Whereas
BY C. F. A. PANTIN, MA.
(From the Zoological Laboratory, Cambridge, and the
Marine Biological Laboratory, Plymouth.)
WHEREAS it is well known that most organisms cannot survive sudden large changes
in the salinity of the external medium yet estuarine organisms have developed
powers of resistance to them. Recent work, particularly that of Schlieper (1929),
has thrown some light on the behaviour of estuarine animals in diluted sea water.
In most estuarine animals the salinity variations of the environment fall far
short of the change from completely fresh water to undiluted sea water. But the
triclad worm, Gunda ulvae, is found where variations of this magnitude may be
expected with each tide, for it inhabits the sea shore at the estuaries of very small
streams.
The ability of these worms to withstand these changes is the more remarkable
because of their small size, and because they are not covered with a protecting
cuticle which might render them impervious to environmental osmotic changes.
Moreover, whereas coelomates may mitigate the effect of such changes upon their
cells by control of the composition of their body fluids, Gunda possesses no body
cavity.
Salinity variations may be expected to influence greatly the water exchange
of Gunda, and experiments showed that this was true (Weil and Pantin, 1931;
Pan tin, 1931). But it was soon found that in order to interpret these experiments
an accurate description of the environment was required.
Little attention has yet been paid to the immediate physical and chemical
environment in which an estuarine organism lives. Thus, although Gunda lives
definitely in association with fresh water, it seems possible that by suitable changes
of position it might avoid gross changes in salinity. Reid (1930) found that in a
small stream flowing over a sandy shore water deep in the sand may retain a fairly
high salinity at low tide. It is therefore necessary to determine the exact distribu-
tion of the Gunda and the salinity variations in its neighbourhood.
64 C. F. A. PANTIN
Further, experiments on the salt exchange of these worms when exposed
fresh water showed that the chemical composition of this water was a factor of
prime importance. The mere description of salinity changes is inadequate: the
problem of estuarine existence is not one of simple dilution but of dilution accom-
panied by changing chemical composition, a factor which may vary in every river.
Indeed, when speaking of estuarine conditions, the term "fresh water" is far too
wide, .and may cover waters which differ among themselves almost as much as
any of them do from sea water. It was therefore necessary to determine the com-
position of the water flowing by the Gunda in their natural habitat.
THE MEDIUM.
Gunda ulvae is found in large numbers in the estuary of a small stream at
Wembury (South Devon) where this flows over an open foreshore. It is not found
except in association with both fresh water and sea water. The external medium
consists entirely of mixtures of stream water with Atlantic sea water.
The following methods were used to analyse the stream water. A large sample
of stream water was collected on July 23rd, 1930, and the following constituents
determined:
(1) Total solids, by evaporation to dryness on a water bath, followed by heating
to 1800 C. to constant weight.
(2) Carbonates, by titration while boiling with H2SO4 to/>H 7, using phenol red.
(3) Chlorides, by titration by Mohr's method with AgNO3 (Treadwell and Hall,
1928).
(4) Sulphates, determined as BaSO4 using the turbidimetric method of Thresh
and Beale (1925).
(5) Calcium, by conversion to oxalate and titration with KMnO4 (Treadwell
and Hall, 1928).
(6) Magnesium, determined as phosphate in the nitrates from (5) using the
turbidimetric method of Thresh and Beale (1925).
(7) Sodium andpotassium, by removal of SO4, Mg and Ca; conversion to chloride,
and the dry weight compared with the AgNO3 titre. The method estimates accurately
the total alkali metals present but does not give accurately the proportion of Na
to K. The exact determination of this was not considered necessary.
Four sets of independent determinations of each constituent were made. The
maximal and minimal values found'are shown in Table I.
The stream water is a typical hard water containing in addition to CaCO3
a large amount of Mg and SO4. The high alkali chlorides probably result from the
cultivation of the land through which it flows. In winter the Cl' rises and the
CO 3 " falls (column 4). The change in salt concentrations is, however, far smaller
than the change in volume of water in the stream.
The other component of the mixture in the estuary is Atlantic sea water, the
composition of which is shown in Table I. This was calculated from Dittmar's
(1884) analysis.
The Adaptation of Gunda ulvae to Salinity
Table I.
Milligrammes per litre.
*H 78 — 77 82
HABITAT.
Gunda is found under stones in the estuary of the Wembury stream. It is rarely
found beneath stones less than 15 cm. in diameter, probably because these are
subject to violent movement in rough weather.
The stream itself rises in the Staddon grits and flows over Middle and Lower
Devonian slates and grits. The basin is about 2 miles long and contains much
cultivated ground. The stream flows rapidly and varies considerably in volume
according to the season. In July 1930 it was roughly 1 to \\ m. in width and
10-15 c m - deep at the mouth. It may reach many times this volume in winter.
It debouches into a fresh-water pool (about 5 m. x 25 cm. deep) at the top of
a beach of shingle (Fig. 1) in Wembury Bay. Beyond the pool is a shelf of shingle
thrown up by the highest tides. The shingle consists of stones averaging 3-5 mm.
in diameter, together with larger stones and boulders. The stream cuts through this
" beach shelf" and flows rapidly down to the edge of the shingle (Station E, Fig. 1).
In the next section (Stations E-M) it flows through channels and pools in the
rocks with some fine shingle and large stones in its bed. Beyond Station M (Fig. 1)
the stream flows between rocks and sand.
The bed of the stream in the middle section of the estuary (Stations E-M) is
fairly constant. But the course through the shingle varies greatly in rough weather,
and much of the water in this region flows through the shingle and not on the
surface. Thus, from July 15th to 19th, 1930, the stream flowed from the pool
over the shingle, decreasing to about one-half of the volume on the land above the
pool; reappearing full-size round Station E. The sea was calm and the bed of the
stream fairly constant. From July 28th till August 1st the stream disappeared into
shingle at the end of the pool, reappearing as several small springs 10-20 m. down
the beach. The sea was then rough, and the shingle altered each tide.
JEB-VIIli S
66 C. F. A. P A N T I N
Observations were made in the fresh-water pool, P, and at Stations A-]VH
extending down the estuary to a point beyond the occurrence of Gunda. The levels
and distances of the more important stations were surveyed for me by Mr M. A.
Spender. These are shown in the vertical section in Fig. i, together with the
heights of tides. T h e latter are based on the Devonport Chart Datum for tides in
the Hamoaze. The observed tides at Wembury agreed well with these except that,
when the sea was rough on July 28th, 1930, large breakers swept above the level of
high-water spring tides and entered the fresh-water pool.
Fresh-water pool
Desert region
40 50 60 80 90 100 1 10
Metres below pool
/•» /"» r\ -Shingle (measured along bed of stream)
/ / / / / « Bock + boulders
Fig. i. Vertical section along estuary of stream, showing substratum and limits of organisms, etc.
At spring tides, July 28th, 1930, fresh-water fauna pushed back to pool, and upper limit of Gunda
at Station D.
The fauna showed a regular sequence from a very rich fresh-water fauna in the
pool to a normal marine shore fauna. The following collections made are typical.
FRESH-WATER POOL P.
Platyhelminth.es Polycelis cornuta1
Hirudinea Glossosiphonia heteroclita
Herpobdella atomaria
Gastropoda Rissoa ventrosa
Crustacea Gammarus pulex
Insecta Larvae of Ephemerida; Plecoptera;
Chironomidae; Trichoptera
1
This organism agrees perfectly with the description (Whitehead, H., 1921), although it has
been suggested that it is restricted in habitat to high land in a manner similar to Planaria alpina
(Hubault, 1927).
The Adaptation of Gunda ulvae to Salinity
STATION B.
Nil.
STATIONS C AND D.
Platyhelminthes Gunda ulvae
Unidentified Acoelan
Archiannelida Protodrilus flavocapitatus
STATIONS E, F, G AND H.
Platyhelminthes Gunda ulvae
Archiannelida Protodrilus flavocapitatus
Crustacea Gammarus sp. (undescribed)
Jaera marina
STATION K.
Coelenterata Actinia equina (one small specimen)
Platyhelminthes Gunda ulvae
Archiannelida Protodrilus flavocapitatus
Gastropoda Patella vulgata (very few)
Crustacea Gammarus sp. (undescribed)
Melita palmata
Jaera marina
STATION L.
Coelenterata Actinia equina
Platyhelminthes Gunda ulvae (one specimen found on
one occasion only)
Gastropoda Craspedochilus cinereus
Patella vulgata
Crustacea Gammarus sp. (undescribed)
Melita palmata
Jaera marina
Sphaeroma serratum
Carcinus maenas
Vertebrata Onos mustelus
STATION M.
As Station L; no Gunda found on any occasion.
SALINITY.
The salinity changes of the environment were determined by measurement of
the electric conductivity of the medium. Samples were taken over the region of the
distribution of Gunda. Their resistance was compared with artificial dilutions made
from one particular sample of sea water taken well clear of the stream mixed with
stream water taken about 300 yards inland. In comparison with this sample of
sea water English Channel sea water taken several miles from the coast had a
conductivity of 103 per cent. Since the water of the stream contains a large pro-
portion of salts it has a fairly high conductivity, approximately equal to that
of a 0 7 per cent, solution of sea water in distilled water. With proper temperature
control the presence of o-oi per cent, of sea water in the river water can be
measured.
All measurements were made at 16-5° C.
The following observations show that the water in immediate contact with the
Gunda must vary from undiluted sea water to water in which the amount of sea
70 C. F. A. PANTIN
water is negligible in comparison with the amount of salts already in the fresh
water.
July 15th, 1930.
STATION E.
Low tide: 3.30 p.m. Sea water (%
3.30 p.m. Surface water 0-03
Under stone with Gunda ... 0-03
12 cm. below stone in shingle 0-07
High tide: 10.0 a.m.
12.20 p.m. (Station under 50 cm. of sea water
and 3-4 m. from water's edge)
Surface water 100
Under stone with Gunda ... 100
5 cm. below stone in shingle 95
The samples were taken by inserting a pipette under stones beneath which it
was supposed Gunda would be found, the presence of the latter being ascertained
after the sample was taken. The deep samples were taken in order to determine the
presence or absence of any water of very different salinity in the neighbourhood
of the Gunda.
Table II shows the successive changes in salinity during a single tide at six
stations down the estuary.
It is evident that the range covers enormous salinity variations. At the upper
end the worms are subject to a small percentage of sea water for perhaps an hour
each tide. At the lower end of the range they are under normal sea water for some
7 hours, and during the remaining 5 hours the sea water may fall to about 10 per
cent, of the normal concentration. In the maximal part of the range the salinity
varies from almost completely fresh water to undiluted sea water with each
tide.
It is apparent that Gunda is normally called upon to make full use of any
powers it has to withstand salinity changes. Indeed its occurrence becomes rare
as soon as these changes become reduced in magnitude (e.g. at Station L, just below
low-water neaps). This is remarkable because Gunda can be kept alive for months
in undiluted sea water, though experiments indicate (Pantin, 1931) that these con-
ditions impair for a time their ability to withstand osmotic changes.
The general problem of adaptation to estuarine conditions is, therefore, pre-
sented in a most striking form by Gunda ulvae. A study of the physiological
mechanisms concerned in such a case as this may be expected to yield more informa-
tion as to the means by which marine forms during evolution become adapted to
fresh-water existence than may be gleaned from most estuarine organisms, which
never come in contact with purely fresh water. Gunda can behave temporarily
as a fresh-water organism: why is it still sufficiently dependent upon sea water to
prevent its range extending into a purely fresh-water environment? In the suc-
ceeding papers an attempt is made to analyse the physiological changes induced
in the worms on transference from marine to fresh-water conditions.
The Adaptation of Gunda ulvae to Salinity
Table II.
High tide: 11.50 a.m.; 12.0 midnight. Low tide: 6.0 p.m. July 19th, 1930.
SUMMARY.
1. The environment of the triclad Gunda ulvae has been studied. This organism
lives on the sea shore in the estuaries of very small streams.
2. The components of the external medium are (a) stream water, which is rich
in Ca and C0 3 , and (b) Atlantic sea water. These are mixed in different proportions
in different parts of the estuary.
3. An analysis of the stream water is given.
4. The habitat of the organism is described. This extends roughly from high-
water neap tides to low-water neap tides. A faunistic survey is given.
72 C. F. A. PANTIN
5. The conditions which control the limits of the habitat of Gunda are discussed.
Between the upper limit of occurrence of Gunda and the place of occurrence of
fresh-water forms there is a region devoid of fauna. This region corresponds roughly
with the span between high-water neap tides and high-water spring tides.
6. Salinity determinations have been made on samples taken from the actual
places where Gunda occurred. It is shown that Gunda has to withstand changes from
completely fresh to undiluted sea water. It may normally be exposed to either
extreme for several hours.
7. Salinity determinations made continually throughout the range of Gunda
show that its environment may vary from one in which it is subjected to the action
of sea water for only about 1 hour at high tide to one in which the sea water is only
diluted to about 10 per cent, of its normal strength for a few hours during low tide.
REFERENCES.
DITTMAR, W. (1884). Challenger Report, Physics and Chemistry, 1, 203.
HUBAULT, E. (1927). Inverttbre's Torrenticoles. Paris: Dulau & Co.
PANTIN, C. F. A. (1931). Journ. Exp. Biol. 8, 82.
REID, D. M. (1930). Journ. Marine Biol. Assoc. 16, 609.
SCHLIEPER, C. (1929). Zeit.f. vergleich. Physiol. 9, 478.
THRESH, J. C. and BEALE, J. F. (1925). The Examination of Water and Water Supplies. 3rd ed.
London: J. and A. Churchill.
TREADWELL, F. P. and HALL, W. T. (1928). Analytical Chemistry, 2, 7th ed. John Wiley and Sons,
Inc.
WEIL, E. and PANTIN, C. F. A. (1931). Journ. Exp. Biol. 8, 73.
WHITEHEAD, H. (1921). Essex Naturalist, 20, 1.